FOODS of the HAWAIIAN GOOSE by PAUL H
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1OS Vol. 49 FOODS OF THE HAWAIIAN GOOSE By PAUL H. BALDWIN In a previous paper (Baldwin, 1945) the distribution and reduction in numbers of the Hawaiian Goose, or Nene (Nesochen sandvicensis), were described. In the present study, the food habits are considered as a step in the analysis of causesof the reduction of the population. Becauseof the practicability of obtaining material, this work is based mainly on the analysis of droppings. We should attempt to learn whether or not food resourcesare a limiting factor in the successof the goosein its environment of today. Former writers have reported that seasonal changes in availability of food in the range of the Nene tend to cause it to shift about between uplands and lowlands at various seasonsof the year. Also, avail- ability of foods has been considered as decisive in influencing the area chosen for nesting. The Hawaiian Goose is confined to the Island of Hawaii and inhabits barren lowland country from sea level to 3000 feet and upland slopes of the mountains, Mauna Loa and Hualalai, from 3000 to 9000 feet. Much of the range consists of barren lava flows which support only a scanty growth of herbs, ferns and shrubs and are entirely devoid of water. Other parts of the range include moist grasslands and open forests. The yearly rainfall in these areas falls between the probable limits of 15 and 120 inches. The low- land range is on the leeward side of the island, hence is hot and dry, whereas the upland range is more variable in climate. The gooseis a vegetarian, utilizing grasses,berries and herbs. The following plants have been mentioned by various authors as providing food: grasses,Fragaria chiloensis (white strawberry), Rubus hawaziensisand R. Macraei (akala), Vaccinium reticulaturn (ohelo) , Styphelia Tameiameiae (pukiawe) , Solanum nodiflorum (popolo) , Coprosma evno&oides (kukainene), and Sonchus oleraceus (sow thistle). It was agreed that the berries of Vaccitium are one of the most important foods. Solanum and Sonchus were also considered important. The Nene went to the lower elevations in winter and spring to take advantage of the new growth of greens for rearing their young, according to Henshaw (1902: 105). They returned to the uplands in summer where they fed on berries. Foods said to be abundant in the uplands in summer were grasses,Vaccinium, and other berries. These were less plentiful during the cold winter months. Abundant foods in the lowlands in winter were grasses,Son&us, and other herbs. In spite of the scanty crop of berries in the uplands during winter, Peale ( 1848 : 250) found Nene subsisting on them in upland lava fields in November. In spring, grassesand Sonchuswere said to continue abundant in the lowlands, and in late spring wild strawberries were obtainable in the mountains. In summer, strawberries continued to bear but went out of season around the end of July on Hualalai (Brigham, 1909: 12): In the fall, berries of sorts were available. Competition from grazing animals was thought by Pope ( 1932: 110) to have been responsible for migration to the lowlands in winter, while Perkins pointed out that cattle destroy the strawberry plants in Nene range. Lamb ( 1937) suggested that pheasants are competitors of the Nene for foods. Concerning differences between the food habits of the Nene and other species of geese, Miller ( 1937 :3) points out that in comparison with winter flocks of geese in North America the Nene probably does less feeding on short grass and on grain growing on open level surfaces and appears to do no foraging in the mud at the edges of ponds and marshes. May, 1947 FOODS OF THE HAWAIIAN GOOSE 109 The knowledge available as summarized above affords a meagre concept of the foods eaten. We find no detailed list of foods used, no mention of the chemical composition of foods or ecological aspects of the food supply. Nothing is said about the nutritional requirements of the geesethemselves. In this study we attempt to supply some of this information, although the subject of the nutritional requirements of the geese could not be included. Procedures.---Clf 640 droppings collected, 543 were microscopically dissected. The material was gathered over a period of six years, between 1938 and 1944. In the field the droppings were readily recognizable. Except for some distinctly small droppings, presumably from juvenal birds, they were fairly uniform in size, and all were typically laminated at right angles to the long axis. A J-shape was characteristic. Mongoose drop- pings were similar in size but not in other features. Pheasant droppings were smaller, not laminated, and not frequently seen. The material varied from bleached to fresh. Volumetric measurements of identified components were made by the water dis- placement method and by visual estimation. It proved possible to identify about 40 per cent of the material. Plant names used in the study are after Fagerlund and Mitchell (1944), and considerable data on the distribution of plants in Hawaii National Park were drawn from this work. The project was carried out on behalf of the United States National Park Service. The University of Hawaii kindly loaned a microscope for the work. LISTOFFOODS The following list of foods includes those actually identified in this study together with those attributed to the diet of the Nene by previous authors. A summary of foods actually found with data on the number of recorded occurrences, the per cent of occur- rences, and the per cent of the total volume for each food is presented in table 1. Agrostis avemcea. Grass. Availability: Frequent along roadsides, Hawaii National Park (henceforth H. N. P.) . Use: Found in droppings from Kau Desert. Seeds, leaves, stems utilized. Axonopus afitis. Narrow-leaved carpet-grass. Availability: Abundant on grazed lands in Humuula (5800 ft.) ; abundant in moist pastures below 2000 ft. (Whitney, Hosaka, and Ripperton, 1939:26). Use: Found in droppings from Humuula. Seeds, leaves, stems utilized. Bidens. Availability: Various species found from sea level to 7ooO ft. on Mauna Loa in H. N. P.; greatest abundance in rather dry areas. Use: Found in droppings from Mauna Loa in H. N. P. and Puuwaawaa. Seeds, leaves, stems utilized. Probably important in lowland areas. Bulbostylis capdllaris. Sedge. Availability: Frequent on volcanic ash deposits in Kau Desert between 2ooO and 4000 ft. Use: Found in droppings from Kau Desert. A minute plant. Mainly seeds utilized. Carex Maclovbm. Sedge. Availability: Frequent on slopes of Mauna Loa in H. N. P., from 4000 to 7000 ft. Use: Found in droppings from Mauna Loa in H. N. P. and Humuula. Seeds utilized and probably . the leaves also were frequently taken. Curex wahuensis. Sedge. Availability: Frequent in moderately wet areas in H. N. P. and occurring up to 10,000 feet on Mauna Loa ; seen at summit of Hualalai. Use: Found in droppings from Mauna Loa in H. N. P., Humuula, and summit of Hualalai. Seeds recovered and also green parts. Is one of the most important foods. Carex. Sedge. Use: Found in droppings from Humuula. Probably is C. wahuensis, but could not be specifically identified. Cirsium vulgare. Common thistle. 110 THE CONDOR Vol. 49 Availability: Frequent in dry to moderately dry areas from the Kau Desert at ~OOCIto 6ooO ft. on Mauna Loa in H. N. P.; found at summit of Hualalai and Puuwaawaa; where rainfall is less than 60 inches, at low and medium elevations. Use: Found in only one dropping from Puuwaawaa. Seed feathers found. An important food. CoProsma wnodeoides, var. typica. Kukainene. Availability: Frequent in moderately wet areas from Kilauea up to about m ft. on Mauna Loa in H. N. P.; abundant in Humuula.; found at summit of Hualalai but not at lower elevations on Puuwaawaa where droppings were collected. Use: Found in droppings from Mauna Loa in H. N. P. and Humuula. Fruits utilized. Perhaps not so important as previously thought. Cyperus polystachyos. Sedge, kilioopu. Availability: Abundant along roads and trails at Kilauea; frequent in Kau Desert; common in low altitude and occasional in medium altitude zones in Hawaii where rainfall is over 60 inches per year (Ripperton and Hosaka, 1942 :4g). Use: Found only in droppings from the Kau Desert. Seeds and green parts utilized. Desckampsia nubigemz. Grass. Availability: Frequent along roadsides about Kilauea and dominant on Mauna Loa slopes from 4000 to 7ooO ft.; seen around 8000 ft. on Hualalai. Use: Found in droppings from Mauna Loa (H. N. P., Humuula.) and summit of .Hualalai. Seeds, leaves, stems utilized. An important food. Digitaria pruriens. Crabgrass. Availability: Infrequent in moderately dry areas in H. N. P. : rather abundant on all the Ha.waiian Islands in wet regions of the lowlands (Whitney, Hosaka, and Ripperton, 1939:54). Use: Found in droppings from Kau Desert and abundantly in droppings from Puuwaawaa. Seeds and green parts utilized. LX&aria G&SC,. Crabgrass. Availability: Frequent in moderately dry areas in H. N. P. ; fairly abundant in moist and semi- dry regions to an altitude of So00 ft. or more (Whitney, Hosaka, and Ripperton, 1939357). Use: Found in droppings from Kau Desert (abundantly) and from Mauna Loa in H.N.P., Puuwaawaa, and Humuula. Seeds especially recovered, but green parts also found. An important food. Festucu m.egaZura.Foxtail fescue. Availability: Of restricted occurrence in H. N. P. around 4000 ft.; found occasionally at medium altitudes in moist situations (Whitney, Hosaka, and Ripperton, 1939: 72). Use: Found in droppings from Mauna Loa in H. N. P. Seeds and green parts utilized. Fragnria chiloetis. White strawberry. AvailaMlity: Frequent in moderately wet forests of medium elevation; in H. N. P. found from perhaps 3ooO to 6000 ft., abundant around 4000. Use: Not recovered from droppings.